Ruminal Metabolism and Intestinal Digestion of Fatty Acids.

William Chalupa, Peter Moate and Ray Boston School of Veterinary Medicine University of Pennsylvania Kennett Square Pa 19348 Summary A lipid sub-model is used to illustrate ruminal metabolism and intestinal digestion of long chain fatty acids (LCFA). Ruminal lipolysis (defined as enzymatic cleavage of ester linkages and dissociation of salts of fatty acids) varies between feeds. Megalac (calcium salts of palm oil fatty acid distillate) was the only ingredient that showed appreciable protection against lipolysis. Lipolysis is a pre-requisite for ruminal biohydrogenation. Polyunsaturated fatty acids, especially C18:2, appear to have an energy independent effect on improving reproduction. However, less than 10% of the C18:3, C18:2 and C16:1 that are in the form of free fatty acids will escape biohydrogenation. C18:1t, which can inhibit mammary synthesis of fat, accumulates in the rumen when there is incomplete biohydrogenation to C18:0. De novo synthesis of fatty acids occurs, but as fatty acid intake increases, the extent of de novo synthesis declines as a result of increased uptake of LCFA by microbes. There is insufficient data to model the effect of rumen active fat on rumen digestion and fermentation so we suggest using the advisory of Jenkins (1997), which is based on unsaturation of LCFA and ration fiber, for the upper limit. Most of the LCFA flowing to the small intestine are free fatty acids but there are also non-lipolysed fatty acids in the form of glycerides and calcium salts and fatty acids in bacteria. In general, intestinal digestion of free fatty acids and non-lipolysed fatty acids in forages, grains, proteins, whole cottonseed, and cracked or ground soybeans and other oil-seeds are similar. Digestion of non-lipolysed fatty acids in tallow, hydrogenated tallow, grease, vegetable oils, animal/vegetable blends, whole-intact soybeans and other whole-intact oil-seeds with the exception of cottonseed is less than digestion of free fatty acids. In particular, digestion of non-lipolysed C18:0 in some feeds is zero. Digestion of non-lipolysed fatty acids in the form of calcium salts is greater than digestion of free fatty acids. To increase absorbed amounts of LCFA like C18:2 and C18:3, they must either be in a form that protects them from lipolysis or large amounts must be fed. The latter, however, can lead to accumulation of C18:1t that can decrease mammary synthesis of fat.

Introduction Fats provide more than calories! They provide specific polyunsaturated fatty acids that participate in a host of metabolic reactions that can have an impact on dairy cow metabolism (Jenkins, 2002; Sanchez and Block, 2001). This is the most exciting

In dealing with this problem. absorbed fat is 95% of the ether extract entering the small intestine. 2000). a new lipid sub-model (Moate et al. Instead. they consider dietary lipid as a single entity. Major issues of the fat sub model include: 1. Dairy cows were fed a diverse range of feeds with a wide range in intakes (mean. CNCPS (Fox et al. Acronyms used in the fat sub-model are defined in Table 1. 2000a. and assuming that LCFA = 0 if EE is less than 1. it has been evident that dairy cow nutrition and nutrient management models are vital to the continued success of the dairy industry. CPM-Dairy and CNCPS operate on ether extract and not fatty acids. 2. We have computer programs like CPM-Dairy (Boston et al. C14:0. 2000) and NRC-Dairy (2001) that allow us to balance rations on the basis of amino acids but no dairy nutrition model describes fat correctly. In addition. C16:1. 6. Intestinal digestibility is also handled is a simplistic manner. 878 g. NRC-Dairy assumes that for diets containing 3% or less ether extract. range. they ignore the transformation processes that affect dietary fatty acids in the rumen and they generally have a very simplistic treatment of de novo production of fat within the rumen. 4. For example. C18:2. C18:1trans. intake of fatty acids ruminal lipolysis of dietary fats biohydrogenation of fatty acids in the rumen de novo production of fatty acids in the rumen effects of fat on rumen digestion and fermentation intestinal digestion of fatty acids Data used to develop this model came from 8 published experiments that reported intakes and flows of LCFA to the duodenum and to feces. NRC-Dairy (2001) advocates measuring LCFA content of feeds or using the equation: LCFA = EE-1 (Allen. 2000). Intake of Fatty Acids Fatty acids and not ether extract are the nutritional entities of importance.development in ruminant nutrition since we recognized that proteins provide amino acids. No model considers specific fatty acids. Dairy-NRC (2001) estimates fatty acids empirically from ether extract. 3. 197 to 1339 g) of LCFA. the digestibility of total LCFA is 100% In this report. . Data in Figure 1 shows that the Allen (2000) equation predicts the LCFA content of some feeds well but it may not be accurate for lush grasses and legumes that contain high amounts of pigment that are ether extract. C18:3).b. In CPM-Dairy and CNCPS. C16:0. However. 5. C18:0. For some years now. 2001) is used to illustrate ruminal metabolism and digestion of LCFA. C18:1cis. The NRC-Dairy model employs different digestion coefficients for total LCFA from different sources and discounts digestibility as dry matter intake increases above maintenance.c. These experiments contained 27 common dietary ingredients in 36 diets. The fat model focuses on the major LCFA (C12:0.

The approach of Waldo et al. CPM-Dairy and CNCPS employ the model of Waldo et al. and C:18:2) account for 20% or more of the total LCFA. As feed intake increases. We use the term “lipolysis” to refer to the liberation in the rumen of LCFA in feed ingredients. There is evidence that. (1972) to define rumen digestibility (RD) as the specific rate (%/h) of rumen digestion (Kd) divided by the specific rate (%/h) of disappearance due to digestion and passage (Kd + Kp): RD =    Kd     Kd + Kp  Variable passage rates provide a method for estimating variations in ruminal digestibility as feed intake changes. We take . the rates of lipolysis may depend upon the concentration of LCFA in the rumen (Beam et al. C18:0.”  Klip   RFLCFA = DIETLCFA *     Klip + Kp  Where RFLCFA (g/d) is the amount of fatty acids in feeds that are lipolysed or converted to a free form in the rumen. We are assembling a bank of feeds that will be analyzed for fatty acids and ether extract by Dr. Klip (%/h) is the rate of lipolysis and Kp (%/h) is the rate of passage. There are over 200 references that contain information on fatty acid composition of feeds but this only includes about 40 different feeds. Lipolysis of dietary fat. by digestion or by passage. The following equation calculates the amounts of LCFA in feeds that are “lipolysed. rate of lipolysis (Klip) and rates of biohydrogenation (Kb) of individual fatty acids replace Kd in the above equation. 2000). Tom Jenkins at Clemson University. When the major fatty acids in feed ingredients (C16:0. Ruminal Lipolysis and Biohydrogenation Nutrients entering the rumen can only disappear from the rumen by two routes. 2001). galactosylglycerides and sterol esters and the dissociation of calcium salts of fatty acids. phospholipids.Many references list fatty acid profiles but not total LCFA. (1972) was extended to describe the extent of lipolysis. We define fatty acids arising from lipolysis as rumen free long chain fatty acids (RFLCFA) and fatty acids that were not lipolysed as rumen non-lipolysed long chain fatty acids (RNLCFA). In this model. DIETLCFA (g/d) is the intake of fatty acids. 1992). This includes enzymatic hydrolysis of acylester linkages in triacylglycerols. rates of passage increase and the extent of ruminal digestion is reduced (Sniffen et al. coefficients of variation are usually less than 20% (Moate. for some lipids (tallow in particular).

C16:1→ C16:0. Kb (%/h) is the rate of biohydrogenation and Kp (%/h) is the rate of passage. the ruminal passage rates of RFLCFA are the same as the feed ingredients from which they were derived. Biohydrogenation is a complex process that involves the formation of many isomers (Jenkins. (1972) was used to calculate biohydrogenation of unsaturated RFLCFA. The lipolysis rate of tallow decreased as the amount of tallow in the ration increased but even at a level of 5% of ration dry matter. At each step. Megalac (calcium salts of palm oil fatty acid distillate) was the only ingredient that showed appreciable resistance to lipolysis (53%). there is opportunity for the specific fatty acid to either pass out of the rumen or to be further biohydrogenated. We adopted a simplified depiction of biohydrogenation of the major unsaturated RFLCFA that describes the essential features of these pathways. Rates of ruminal lipolysis vary depending on the feed ingredient (Table 2). Unsaturated LCFA not liberated by lipolysis escape biohydrogenation. After examining the flows of specific LCFA to the duodenum.account of this by using moderating factors to allow the Klip to be adjusted in response to different levels of total fatty acids from specific feeds in the total diet: Klip = K * Exp(-L * DIETLCFA) Where Klip (%/h) is the rate of lipolysis adjusted for LCFA in feeds. L is a lipolysis adjustment factor for the affect of DIETLCFA on lipolysis and DIETLCFA is the percentage of the total diet that is LCFA from a specific feed. RFLCFA is the amount (g/d) of specific free fatty acids that are produced by lipolysis. RFLCFA are biohydrogenated in stepwise processes: C18:3 → C18:2 → C18:1t → C18:0. Most feeds have high rates of lipolysis and are therefore extensively lipolysed in the rumen. only rumen free unsaturated fatty acids are biohydrogenated. The model considers the biohydrogenation of the RFLCFA derived from each feed separately but we assume that the biohydrogenation rates (Kb) are independent of the feeds from which the RFLCFA were derived. K (%/h) is the maximum rate of lipolysis.  Kb  BHLCFA = RFLCFA *   Kb + Kp     Where BHLCFA is the amount (g/d) of specific fatty acids that are produced by biohydrogenation. In our model. we observed that the ‘estimated concentration’ of RFLCFA appeared to influence the rates of . The model assumes that lipolysis is a prerequisite for biohydrogenation. 2002). C18:1c → C18:0. Klip = K. In this model. Thus. 92% of tallow fatty acids are lipolysed. Biohydrogenation. A model based on that of Waldo et al. When L = 0.

B * RFLCFA  KbRFLCFA = K b* EXP   DMI   Where KbRFLCFA (%/h) is the adjusted rate of biohydrogenation. However. Kb (%/h) is the theoretical maximum rate of biohydrogenation. In this model we assume that each feed containing fermentable carbohydrate has the potential to induce in the rumen. Feeds contain little C18:1t. Data in Figure 3 present the extent of ruminal biohydrogenation and. He concluded that the flow of LCFA to the duodenum is generally closely related to. more than 90% of the rumen free C18:3. The amount of rumen free LCFA affected the biohydrogenation rates of C18:3. B is the adjustment factor for the effect of RFLCFA on biohydrogenation. In order to take into account this negative effect of fatty acids on de novo . It is interesting that as the level of rumen free C18:1t increases. RFLCFA (g/d) is the amount of rumen free LCFA and DMI (kg/d) is dry matter intake. C16:0. by difference. Rates of biohydrogenation are in Figure 2. De Novo Production of LCFA in the Rumen Jenkins (1993) reviewed the literature on the factors affecting the balance of LCFA across the rumen of sheep and cattle.biohydrogenation. but is usually slightly higher than the dietary LCFA intake. Jenkins. Therefore we again employed exponential moderating equations to allow for adjustment of biohydrogenation rates:  . 2000. 2002). appear to have an energy independent effect on improving reproduction in the dairy cow. de novo production of LCFA. There are considerable differences in rates with C18:3>C16:1>C18:2>C18:1t>C18:1c. De novo synthesis of fatty acids occurs. through growth of ruminal bacteria. The main LCFA that are produced de novo are C18:0. the extent of de novo synthesis may decline as a result of enhanced uptake of exogenous LCFA by microbial cells. but at high Fatty acid intakes. especially C18:2. Mammary uptake of C18:1t appears to inhibit synthesis of milk fat (Chalupa and Sniffen. biohydrogenation decreases so that more C18:1t will flow to the small intestine. Because the rates and extents of biohydrogenation of C18:3 and C18:2 are greater than the rate and extent of biohydrogenation of C18:1t. it is easy to see how there will be increased absorption of C18:1t when diets contain polyunsaturated fatty acids in forms that have high rates of lipolysis. It is produced through biohydrogenation of C18:3 and C18:2. C18:2 and C16:1 will be biohydrogenated so that 10% or less will escape the rumen. the percentage of rumen free unsaturated fatty acids that will escape biohydrogenation and pass to the small intestine. C16:1 and C18:1t but had no affect on biohydrogenation rates of C18:2 or C18:1c. Polyunsaturated fatty acids. C16:1 and COther.

LCFAUP is the adjustment factor for the uptake of RFLCFA by microbial cells. diets that contained hydrogenated tallow and whole-intact soybeans had lower digestibilities. In general. we again employed exponential moderating factors to adjust the rate of de novo synthesis of LCFA in relation to the amount of the relevant RFLCFA produced in the rumen as a result of lipolysis and biohydrogenation. which is based on unsaturation of fatty acids and ration fiber. there is insufficient data to model the effect of rumen active fat on rumen digestion and fermentation. LCFASYN is the de novo production of specific LCFA per gram of FTCHO. We suggest using the advisory of Jenkins (1997). bacteria take up more LCFA and de novo synthesis decreases. However. as shown in Figure 5. there are strong linear relationships between the flow of LCFA to the duodenum and intestinal absorption. Intestinal Digestion of Fatty Acids Most of the LCFA flowing to the small intestine are in the form of rumen free fatty acids but there are also non-lipolysed fatty acids and fatty acids in bacteria. The generalized formula is  . as the amount of RFLCFA increases. RFLCFA (g/d) is the amount of rumen free LCFA and DMI (kg/d) is dry matter intake. Effects of Fatty acids on Rumen Digestion While we can predict the level of rumen free LCFA in the rumen. Unprotected fat (%DM) = (6*ADF)/UFA or (4*NDF)/UFA Where ADF and NDF are expressed as a percentage of ration DM and UFA are unsaturated fatty acids (C18:1 + C18:2 +C18:3) expressed as a percentage of total fatty acids.LCFAUP * RFLCFA  RPLCFA = LCFASYN*FTCHO * EXP   DMI   Where RPLCFA (g/d) is the de novo production of LCFA. It is likely that the intestinal digestibility of duodenal free fatty acids derived from different ingredients is the same but the digestibility of fatty acids that escaped ruminal lipolysis could differ. The impact of dry matter intake and level of dietary fatty acids is shown in Figure 4. for the upper limit of rumen active fat. However. . As dry matter intake increases there is more FTCHO (and bacterial growth) so that de novo synthesis increases. FTCHO (g/d) is fermentable carbohydrate in feeds.synthesis of LCFA.

479 g. . (3) hydrogenated tallow.98. (1997) and Moller (1988) where the apparent digestibilities of C16:0 and unsaturated C18 fatty acids were elevated in diets containing calcium salts of palm or rapeseed fatty acids. C18:1c. it is apparent that there is close concordance between measured and predicted flows of total LCFA to the duodenum (R2=0. (4) wholeintact soybeans and other whole-intact oil-seeds with the exception of cottonseed. The low correlation (R2=0. the digestion coefficients (Table 3) for RFLCFA and rumen non-lipolysed LCFA in feeds in category 1 are similar. 72 to 1040) than intakes in the data used to develop the model (mean. This is consistent with the findings of Enjalbert et al. the digestion coefficient for C18:0 was 0. Thus it seems that the often reported low digestibility of C18:0 from hydrogenated fat sources may be more likely due to low or inefficient intestinal emulsification than to an ineffective lipase system. whole cottonseed. C18:0. proteins. substantially less than the corresponding coefficient for RFLCFA. Thus. Due to the scarcity of published experiments with the requisite in vivo data. (1992) reported that when cows were fed emulsified vegetable fat protected by means of formaldehyde-casein. 3 and 4 are zero. and cracked or ground soybeans and other oil-seeds.We used optimization techniques to derive digestibilities for RFLCFA (including bacterial fatty acids) and six classes of rumen non-lipolysed LCFA: (1) forages.92. The predicted bias was 13% or less. From data in Figure 6 and Tables 4 and 5. there were only two experiments with lactating Holstein*Friesian dairy cows while the remaining experiments involved steers of various ages and breeds. the digestion coefficients for rumen non-lipolysed C18:0 in categories 2.61) and high bias (75%) between measured and predicted flows of C16:1 probably reflects the low flow (mean flow=2 g/d) of C16:1. Data in Table 4 show that there was a high correlation (R2>0. Model Validation Data used to validate the model came from 8 published experiments that reported intakes and flows to the duodenum and feces of the major LCFA (Moate. B ø rsting et al. C18:2. (2) tallow. range. 197 to 1339 g). In general. 2001). C18:1t.99. (5) fish meal supplements and (6) calcium salts of fatty acids. intakes of LCFA were less (mean. The digestion coefficients for all of the rumen non-lipolysed fatty acids from feed categories 2 to 5 are less than. grease. and in many cases. In particular.91) between measured and predicted flows of C16:0. range. vegetable oils and animal/vegetable blends. bias=5%) and measured and predicted absorption of total LCFA from the intestine (R2=0. C18:3 and COther to the duodenum. bias<1%). In contrast. The digestion coefficients for the major rumen non-lipolysed fatty acids in the form of calcium salts are substantially greater than the coefficients for RFLCFA. grains. 878 g.

The predicted bias was12% of less for C12:0. C18:1t can inhibit mammary synthesis of fat. However. This reflects the intense biohydrogenation of C18:unsaturated free fatty acids in the rumen. When fatty acid supplements are added to rations. Intestinal digestibility of Megalac and Megalac R is higher than the basal diet because rumen non-lipolysed fatty acids in the form of calcium salts have higher intestinal digestibilities than rumen non-lipolysed fatty acids in the form of glycerides. 26% corn silage. 22% steam-flaked corn. The basal diet provided 500 g LCFA (2% DM basis). C18:0. intakes of C18:0 are low but substantial amounts of C18:0 reach the small intestine.7% fat. C18:2. Application of the Model To demonstrate the application of the model. C18:0. It is thus possible that today’s high producing cows may be . 400 g of each fatty acid supplement raised dietary LCFA to 3. C18:1t can accumulate in the rumen. 400 g of LCFA were provided by adjusting the proportions of mineral mix and fatty acid supplement (Table 6). C16:0. Feeding 400 g of LCFA in the form of whole cottonseed and roasted soybeans doubled the amount of C18:t absorbed. On an energy basis. Absorption of C16:1. only small amounts (2 to 3 g/d) of C16:1 and C18:3 were absorbed and COther is a “mixed bag” of LCFA not always reported in all experiments. As noted before. Cow’s milk contains approximately 2 to 6% of the fatty acid content as C18:2 (Sanchez and Elliot. we fed a 650 kg cow 25 kg of a diet that contained (DM basis) 26% alfalfa silage. The basal ration in Table 6 contained 225 g C18:2. C18:2 appears to have an energy independent effect on improving reproduction in the dairy cow. C18:3 and COther was predicted poorly. 2001) so our example cow would secret 30 to 89 g C18:2 in milk. the bias was about 20%.6%. Megalac R. C18:1t. and C18:2. C18:1c. 14% soybean meal. The amounts of C18:1t absorbed from Megalac. only 58 g reached the duodenum with 48 g absorbed. Total long chain fatty acids. 2% blood meal and 10% mineral mix/fatty acid supplement. Energy Booster and tallow are small. bacterial cells will take some of the RFLCFA up with a concomitant decrease in de novo production. C14:0. De novo production of LCFA only occurred on the basal diet because the fatty acid supplements provide no or little fermentable carbohydrate for microbial growth. There is little C18:1t in feeds but because of biohydrogenation of C18:3 and C18:2 to C18:1t and incomplete biohydrogenation of C18:1t to C18:0.86) between measured and predicted absorption of C12:0. C18:1t. With the exception of Energy Booster. C16:0 and C18:0 and C18:2. For C18:t and C18:1c. but because of extensive biohydrogenation in the rumen. the basal ration would support production of 40 kg milk with 3.Data in Table 5 show that there was a high correlation (R2>0. As noted before. C14:0.

and D.. Proc Australian Soc. Dairy Sci. J. 2000. D. and V. C. The net carbohydrate and protein system for evaluating herd nutrition and nutrient excretion. Sect. J. R. T. Sniffen. Chase. Hvelplund. and T.deficient in this essential fatty acid. Dairy Research. ed. Department of Animal Science. P. C. Sci. Enjalbert. J. Beever. J. milk composition and physical properties of butter. M. F. Nicot. E. Moncoulon.. Boston. Weisberg. The conversion of a scientific model describing dairy cow nutrition and production to an industry tool: the CPM Dairy Project. 76:3851-3863. T. Bayourthe. Jenkins. E. Anim. G. Munson and W. M. Mc Namara. Cornell University. 2000. 1997. Kohn and D. Chalupa. M. fish oil or saturated fat. 83:1598-1624. N. P. A. Effects of amount and source of fats on the rates of lipolysis and biohydrogenation of fatty acids in ruminal contents. W and C. Van Amburgh. M. Tedeschi. P. C. Chalupa. Feedstuffs 69(2):14. F. A. To increase amounts absorbed. 64: 181-195. E. Animal Sci.. Lipid metabolism in the rumen. France. dietary C18:3 must either be in a form that protects it from lipolysis or large amounts must be fed to increase absorbed amounts. Palmquist. Janczewski. and R. In ‘Modeling nutrition utilization in farm animals’. C18:3. J. Fox. M. T. Success of fat in dairy rations depends on choosing the right amount.. Model Documentation. G. 2000. Effects of dietary calcium soaps of unsaturated fatty acids on digestion. R. R. C. C. 1993.. J. J. T. L. R. Agric. Ithaca. Bø rsting. 130 Morrison Hall. As with C18:2. 42:148-156. New York 14853-4801. A. Beam. . Acta. Scand. Balancing Rations for Milk Components. R. Jenkins. C. Sniffen. C.2000. Fatty acid digestibility in lactating cows fed increasing amounts of protected vegetable oil. Pell.Animal Science Mimeo 213. N. Dairy Sci. Moate. Tylutki. L. Overton. M. Fox. 83:2564-2573. Sydney. O. Durbal. S. Jenkins. J. T. C18:2 must either be in a form that protects it from ruminal lipolysis (Megalac R) or the feed ingredient must contain high amounts of C18:2 (soybeans). Pp361 – 377. Feeds contain C18:3 but little reaches the intestine because of rapid biohydrogenation in the rumen. Effects of diet on short-term regulation of feed intake by lactating dairy cattle. M.. D. 2000. References Allen. L. 1992. Rasmussen. there is also an increase in absorbed C18:1t which might lower milk fat test. With an ingredient like soybeans. C. Dairy Sci. 1997. however. Vernay.

D. Fox.: National Academy Press. and W. Unpublished data. J. Dairy Sci.1):279. Dairy Sci. J. L. 2000. Ed. 2000.. G. 83(Suppl. R.Jenkins. 1988. Minneapolis.C..J. Proc. 2002. J. Chalupa.J. J.J. Biolinol Futterfette. 23-40. P. Proc.. W. Boston. R. Chalupa. P. W. Boston. Dairy Sci. Chalupa. 83(Suppl. Moate. Dairy Sci. Southwest Nutrition and Management Conf. and W. P. 70:3562-3577. 55:125-29. Van Soest. Ruminal lipolysis and biohydrogenation of long-chain fatty acids. 2001. Waldo. O’Connor.1):352. National Research Council. J. D. P.B. The influence of high amounts of fat or Ca-soaps to dairy cows on intestinal absorption of fatty acids and digestibility of structural carbohydrates. Fatty acids for dairy cows: More than just calories. Moate. In: Futterfette in der Tierernahrung. R. Digestion of Long-chain fatty acids in dairy cows. Moller. Smith. Russell. Pennsylvania. 2001. Ed. Chalupa.1):280. Means and standard deviations of ether extract. Sanchez. P.1):279. Kennett Square. 1972. Moate. P. J. Phoenix. Model of cellulose disappearance from the rumen. Validation of a model for the digestion of fat in dairy cows. 2000. Moate. 83(Suppl.C.J.C.. 2001. R.J.C. T. Caloric versus noncaloric considerations when feeding fat to dairy cattle. . Univ.) Washington D. Ziegelitz. L. total fatty acids and individual fatty acids in common dairy feeds. P. Ruminal production of long-chain fatty acids. D. Sci. 84(Suppl. A net carbohydrate and protein system for evaluating cattle diets: II Carbohydrate and protein availability. and W. R. D. C. and J. Nutrient Requirements of Dairy Cattle (7th Rev. R. Minnesota Nutrit. and W. and E.K and Block. 1992. J. Moate. J.C. Conf. Boston. Sniffen. J. Boston. E. 2001. Cox.Anim. Dairy Sci.

. Relationship between ether extract and total fatty acids. Effect of level of unsaturated rumen free long chain fatty acids (RFLCFA) on the rate of biohydrogenation. 500 450 400 350 300 250 200 150 100 50 0 0 1 2 3 18:3 16:1 100 90 80 70 60 50 40 30 20 10 0 0 1 2 3 Biohydrogenation (%/h) Biohydrogenation (%/h) 18:2 18:1c 18:1t 4 5 4 5 Unsaturated RFLCFA (%DM) Unsaturated RFLCFA (%DM) Figure 2.Total Fatty Acids (%DM) 5 4 3 2 1 0 0 Corn silage Alfalfa hay Corn Grain Soybean meal Grass hay EE-1 1 2 3 4 Ether Extract (%DM) 5 Figure 1.

The dietary concentration of total LCFA was varied by adjusting the proportions of mineral mix and fatty acid supplement.Biohydrogenation (%RFLCFA) 95 90 85 80 75 70 65 60 0 1 2 3 4 5 Rumen Escape (%RFLCFA) 100 40 35 30 25 20 15 10 5 0 0 1 2 3 4 5 Unsaturated RFLCFA (%DM) 18:3 18:2 18:1c 18:1t 16:1 18:3 Unsaturated RFLCFA (%DM) 18:2 18:1c 18:1t 16:1 Figure 3. 26% corn silage. The extent of rumen escape [100 – biohydrogenation]. De Novo Synthesis (g/d) 180 160 140 120 100 80 60 40 20 0 0 2 10 kg DMI 15 kg DMI 20 kg DMI 25 kg DMI 4 6 Total Fatty Acid (%DM) 8 10 Figure 4. 2% blood meal and 10% mineral mix /fatty acid supplement. Effect of level of unsaturated rumen free LCFA (RFLCFA) on the extent of biohydrogenation [(Kb/(Kb+Kp))*100 where Kb is from Figure 2 and Kp = 7 %/h]. 22% steam-flaked corn. Effect of level of dietary fatty acids and dry matter intake on predicted de novo production of total LCFA. The diet contained (DM basis) 26%alfalfa silage. . 14% soybean meal.

7) + 0.1800 Y = 17.99 1200 600 0 0 600 1200 Duodenal Flow of Total LCFA (g/d) 1800 Figure 5. R = 0.02) *DTotalLCFA 2 N = 27.74 (0.7 (16. Measured and predicted duodenal and absorbed total long chain fatty acids. corn and a protein source. Diets containing hydrogenated fat (hollow diamonds) or intact soybeans (hollow squares) were excluded from the regressions. Relationship between duodenal flow and absorbed total LCFA. 1200 1000 800 Data Data Predicted (g/d) Predicted (g/d) residual Y=X 600 400 200 0 residual Y=X 800 600 400 200 0 -200 0 200 400 600 800 1000 1200 0 -200 200 400 600 800 Measured (g/d) Measured (g/d) Absorbed Total LCFA Duodenal Total LCFA Figure 6. diets (dot) contained Megalac. Different symbols represent different diet types: most diets (solid triangles) contained corn silage. .

Table 1. Acronyms. DIETLCFA g/d Intake of LCFA from feeds DMI kg/d Dry matter intake of feeds EE % Ether extract in the dry matter of feed FTCHO g/d Fermentable carbohydrate in feeds K %/h Maximum rate of lipolysis Kb %/h Maximum rate of biohydrogenation Rate of biohydrogenation adjusted for unsaturated RFLCFA KbRFLCFA %/h Kd %/h Rate of rumen digestion Klip %/h Rate of lipolysis of adjusted for LCFA in feeds Kp %/h Rate of passage rate feeds L decimal1 Adjustment factor for the affect of DIETLCFA on lipolysis LCFA g/d Long chain fatty acid LCFA%j % Percentage of the total diet that is LCFA LCFASYN g/d De novo production of specific LCFA per gram of FTCHO LCFAUP g/d Adjustment factor for the uptake of RFLCFA by microbial cells RD decimal1 Rumen digestibility of feeds RFLCFA g/d Free fatty acids that are produced by lipolysis in the rumen RNLCFA g/d Fatty acids that were not lipolysed in the rumen RPLCFA g/d De novo production of LCFA in the rumen . their units and definitions used in the fat sub-model Acronym Unit Definition 1 B decimal adjustment factor for the effect of RFLCFA on biohydrogenation BHLCFA g/d Amount of specific LCFA produced by biohydrogenation C12:0 Lauric acid C14:0 Myristic acid C16:0 Palmitic acid C16:1 Palmitoleic acid C18:0 Stearic acid C18:1c Oleic acid C18:1t Vaccenic acid C18:2 Linoleic acid C18:3 Linolenic acid COther LCFA other than those listed above and with more than 12 carbon atoms.

L is the constant that describes the effect of level of fatty acid from the ingredient on K 5. K is the lipolysis rate (%/h) without correction for the effects of level of fatty acid from the ingredient 4. Number of diets in data set with this feed 3.Table 2. Tallow fatty acids at 0. 1 1 35 0 35 roasted) Fish meal 1 2 23 0 23 Sunflower oil 1 2 52 0 52 Tallow 1 1 500 0.997 98 77 47 1. Klip/( Klip + Kp ) assuming Kp = 7%/h for concentrates and 5%/h for forages 7. raw) Soybean (whole1 1 16 0 16 intact. roasted) Soybean (cracked.0% of dry matter intake . 1 1 26 0 26 roasted) Soybean (ground.37 79-5007 0 392 Animal Vegetable 2 6 392 Hydrogenated fat 1 1 18 0 18 Megalac 3 5 6 0 6 Lipolysis(%) 6 99 99 99 93 64 38 83 98 81 77 99 99 99 81 68 56 70 79 83 77 88 92 .1 to 5. Lipolysis of fat in feed ingredients Feed Expts1 Diets2 K3 L4 Klip5 Corn silage 6 28 500 0 500 Alfalfa silage 3 16 500 0 500 Alfalfa haylage 3 12 479 0 479 Alfalfa hay 1 5 65 0 65 Pasture hay 1 5 9 0 9 Orchardgrass hay 1 3 3 0 3 Corn 1 3 35 0 35 Corn (ground) 6 26 309 0 309 Barley 1 5 29 0 29 Soy hulls 1 4 17 0 17 Cottonseed (whole) 1 2 500 0 500 0 500 Rapeseed (crushed) 1 2 500 0 500 Soybean meal 4 19 500 Soybean (extruded) 2 2 29 0 29 Soy plus 2 8 15 0 15 Soybean (whole1 1 9 0 9 intact. Klip (%/h) = K * Exp (-L * TLCFA%) where TLCFA% is the % of the total diet that is LCFA from the feed ingredient 6. Number of experiments in data set with this feed 2.

95 0.86 1.83 0.64 0 0.73 0. proteins.95 0.72 0.00 0.78 0.67 0 0.19 0.72 0.73 0. grease. and cracked or ground soybeans and other oil-seeds 2.61 0.82 0.79 0. Optimized digestion coefficients for rumen free LCFA (RFLCFA) and non-lipolysed LCFA Category of non-lipolysed LCFA 1 2 3 4 5 6 LCFA RFLCFA Feeds1 Fats2 Hydrogenated Tallow Whole-intact oil-seeds3 Fish Meal Megalac C12:0 C14:0 C16:0 C16:1 C18:0 C18 :1c C18 :1t C18:2 C18:3 Cother 0. Non-lipolysed LCFA from tallow. grains.95 1.95 0.47 0.58 0.16 0 0. whole cottonseed.73 0.79 1.Table 3.67 0 0.83 0.67 0.59 0.18 0 0 0.65 0.83 0.83 0.89 0.56 0.64 0.60 0.95 0.10 0 0 0 0.77 0.78 0.00 0.89 0. vegetable oils and animal/vegetable blends 3. Non-lipolysed LCFA from whole-intact soybeans and other whole-intact oil-seeds with the exception of cottonseed .78 0.73 0.40 0 0.73 0.40 0.75 0.45 0.00 1. Non-lipolysed LCFA from forages.64 0.59 0.59 0.78 0.54 0 0 0 0 0 0 0.

02 0.8 0.46 -18 COther 27 24 10 14.01 0.61 +75 C18:0 36 237 116 37.07 0.89 0.43 +62 C18:0 36 167 83 38.92 -13 1 C18:2 36 27 25 0 0.98 +0<10 C12:0 20 4 8 0.93 +20 C18:1trans 1 C18:1cis 8 25 6 0 00 0.96 0+9 C18:1trans 8 23 14 01 1.99 0+5 C12:0 Flow statistics not calculated because some C14:0 C12:0 and C14:0 are absorbed from the rumen C16:0 36 97 55 01 1.300 0.78 0.05 0.94 0=7 C18:1cis 8 35 14 01 0.14 0.91 0.86 0.92 -22 20 18 0100 0. Measured vs predicted absorption of LCFA from the intestine LCFA n Mean STD Intercept Coefficient R2 Bias (%) 1 Total 36 338 147 0 00 1.87 0.00 0.89 0.13 0.91 -12 C18:2 34 C18:3 31 3 4 1.93 0.95 0-9 C18:3 36 5 7 01 1.91 0=2 COther 27 36 11 01 0.04 0.Table 4.48 0.86 +10 8 15 8 0100 1.86 0. Intercept in original regression was not significantly (P>.87 0.99 0.97 +02 C16:1 16 2 1 1. Intercept in original regression was not significantly (P>.6 0.05) different from 0 so the subsequent regression was forced through 0 Table 5.98 0-7 C14:0 15 C16:0 36 71 44 0100 1.70 0.96 0-1 1.30 0.49 0.16 0-7 1.99 0.33 0. Measured vs predicted flows of LCFA to the duodenum LCFA n Mean STD Intercept Coefficient R2 Bias(%) 1 Total 36 477 206 00 1.99 +10 1 7 4 0 00 0.99 0+5 C16:1 16 2 1 01.05) different from 0 so the subsequent regression was forced through 0 .

6 0.2 Absorbed 48 17 76 0.0 1.2 157 230 18.00 0.5 10.Table 6.9 0.0 2.0 13. Intestinal flows and absorption of LCFA (g/d) predicted by the CPM-Dairy lipid sub-model1 Energy Measurement Basal Megalac Megalac R Booster WCS RSB Tallow Fat Supplement (g) 0 474 474 404 2395 2222 460 Klip (%/h) 6 6 500 500 37 277 Non-lipolysed LCFA (%Intake) 54 54 0 1 16 2 Total long chain fatty acids Intake 500 400 400 400 400 400 400 Duodenum 659 400 400 400 404 404 400 De novo production 159 0 0 0 4 4 0 Absorbed 479 327 337 291 300 298 293 Intestinal Digestion (%) 73 82 84 73 74 74 73 C18:0 Intake 16 16 14 163 10 4 72 Duodenum 330 73 104 201 291 210 212 Absorbed 239 56 78 146 211 153 153 C18:1trans Intake 0.5 0.1 0.9 30.7 12 54 2.8 C18:3 Intake 23.1 1.2 1.00 1.6 0.8 9.0 0.4 C18:2 Intake 225 28 127 7.0 34.0 2. 2% blood meal and 10% mineral mix/fatty acid supplement.3 0.0 0.3 39.8 1. 26% corn silage.1 1.7 5.0 1.6 Duodenum 1.2 4. 14% soybean meal.9 0. .4 9.0 5.8 Duodenum 58 17 77 0.6 10 43 1. 25 kg of a diet containing (DM basis) 26% alfalfa silage. 22% steam-flaked corn. 400 g of fatty acids was provided by adjusting the proportions of mineral mix and fatty acid supplement.0 1.5 23.8 0.9 0.1 0.8 18.3 11.1 Absorbed 1.8 31.6 Absorbed 29.2 Duodenum 37.